Food Hydrocolloids 133 (2022) 107963

Contents lists available at ScienceDirect

Food Hydrocolloids
journal homepage: www.elsevier.com/locate/foodhyd

β-glucans obtained from beer spent yeasts as functional food grade additive:
Focus on biological activity
María Agustina Caruso a, Judith Araceli Piermaria a, Analía Graciela Abraham a, b,
Micaela Medrano a, c, *
a
Centro de Investigación y Desarrollo en Criotecnología de Alimentos- CIDCA (CCT-CONICET La Plata, UNLP, CIC-PBA), 47 y 116, 1900 La Plata, Buenos Aires,
Argentina
b
Área Bioquímica y Control de Alimentos, Facultad de Ciencias Exactas, UNLP, 47 y 115, 1900 La Plata, Buenos Aires, Argentina
c
Instituto Ciencias de la Salud, UNAJ. Av. Calchaqui 6200, B1888, Florencio Varela, Buenos Aires, Argentina

A R T I C L E I N F O A B S T R A C T

Keywords: Nowadays, there is an increasing interest in the search for new healthy compounds obtained from by-products of
Brewers spent yeast the food industry, because of the concomitant trend towards reutilization and by-products exploitation, both
Saccharomyces cerevisiae hallmarks of the circular economy. One of these by-products are the brewers spent yeasts (BSY), generated by the
β-Glucans
brewing industry. BSY, if not properly inactivated and disposed-off, represents an environmental problem. One of
Nutraceuticals
BRM
the uses given to it is as a feed for livestock. However, certain components of yeast cell walls (in particular,
Prebiotic β-glucans) have great potential to be used as Biological Response Modifiers (BRM) for humans. In recent years
the interest on these compounds has grown-up, as the research on its potential application as food additives
capable of conferring a benefit on consumer’s health. Prebiotic, immunomodulatory and antitumoral activities
are particularly of great interest. Since β-glucans possess several physiological functions and considering that BSY
cell walls are rich in β-glucans, the food industry could generate additional revenue by isolating β-glucans from
SBY as a high-value product. The possibility of producing value-added products such as β-glucans from this by-
product could benefit beer manufacturers and, in addition, minimize the environmental impacts from these
industries. This review summarizes the potential application of BSY by-product by the purification of cell wall
components (β-glucans) for application as functional additive, focused on their biological effect on the intestinal
context.

1. Introduction: β-glucans from brewers spent yeasts
β-Glucans are described as polymers of glucose which are widespread
in many bacteria, fungi, algae, and higher plants. In the case of the fungi
kingdom, the presence of β-glucans in cell walls is taxonomically rele­
vant, being its presence characteristic of the Phylum: Oomycota, Chy­
tridiomycota, Ascomycota, Basidiomycota and Deuteromycota
(Alexopoulos, 1962). Among Deuteromycota, Saccharomyces sp. is one of
the most important sources of β-glucans. The cell wall of these yeasts
contains about 55–65% of β-glucan (Klis, Mol, Hellingwerf, & Brul,
2002). The most important source of yeasts is the beer industry; indeed,
in the brewing process, brewer’s spent yeast (BSY) (also known as re­
sidual yeast or surplus yeast) is one of the predominant by-products, and
accounts for approximately 1.5–2.5% of the total beer production.
Global beer production exceeded 1.82 billion hL in 2020 (Web
Reference 1). BSY accounts for a maximum 15% of total by-products
generated during the brewing process (Kerby & Vriesekoop, 2017).
BSY is recovered by sedimentation before full maturation of beer at the
final stage of the fermentation and can be re-used a limited number of
times depending on yeast strain and good manufacturing practices
(Marson, de Castro, Belleville, & Hubinger, 2020).
Because of some inconveniences such as low shelf life, transportation
cost and requirement for further processing, spent yeasts are usually
disposed-off in the environment (Boateng, Okai, Frimpong, & Zeebone,
2015). Nevertheless, the valorisation of BSY can be achieved through the
extraction and isolation of economically important components such as
proteins, amino acids, bioactive compounds like β-glucans and func­
tional peptides, vitamins, minerals, fibre, flavour compounds and
others.
Brewing yeasts can be divided into two classes: top fermenting ale

* Corresponding author. CIDCA (CCT-CONICET La Plata, UNLP, CIC-PBA), 47 y 116, 1900 La Plata, Argentina.
E-mail address: mmedrano@biol.unlp.edu.ar (M. Medrano).

https://doi.org/10.1016/j.foodhyd.2022.107963
Received 11 March 2022; Received in revised form 23 June 2022; Accepted 8 July 2022
Available online 11 July 2022
0268-005X/© 2022 Elsevier Ltd. All rights reserved.
M.A. Caruso et al.
yeasts (Saccharomyces cerevisiae strains) and bottom fermenting lager
yeasts (Saccharomyces pastorianus or S. carlsbergensis strains, a natural
allotetraploid hybrid from S. cerevisiae and S. eubayanus) (Burini, Eiza­
guirre, Loviso, & Libkind, 2021). Top and bottom fermenting brewing
strains have similar surface ultrastructure, but different cell wall elas­
ticity, degree of hydrophobicity, and polysaccharide
structure-composition. The surface of S. pastorianus is poor in proteins
and much more hydrophilic than top fermenting species, explaining
their tendency to sink, contrary to the floating properties of the
S. cerevisiae due to their hydrophobic association with CO2 bubbles
(Alsteens et al., 2008). In ale fermentation, between 1.3 and 1.7 g of
yeasts per litre of beer produced are generated (Marson et al., 2020).
Given the interest in the consumption of foods and additives with
healthy properties, there has also been an increased interest in the
search for additives able to provide a benefit to consumer’s health. In
this sense, β-glucans meet this requirement, and the large number of
scientific papers that have recently been published, which explore
different aspects of β-glucans, demonstrates this. Thus, there are recent
review works that address the application of β-glucans from different
origin (Barsanti, Passarelli, Evangelista, Frassanito, & Gualtieri, 2011;
Cerletti, Esposito, & Iacoviello, 2021; Murphy, Rezoagli, Major, Rowan,
& Laffey, 2020), as well as others that are dedicated to yeast β-glucans.
Within the latter ones, some address towards the use of whole yeasts,
including intracellular content (“yeast extract”) and proteins (Ferreira,
Pinho, Vieira, & Tavarela, 2010; Marson et al., 2020; Puligundla, Mok, &
Park, 2020; Rachwal, Waśko & Polak-Berecka, 2020; Sceni, 2021).
Among the scientific works related to β-glucans isolated from yeasts cell
wall, some of them focus specifically on specific health topics such as
immunomodulatory aspects (Bastos et al., 2022; De Marco Castro,
Calder, & Roche, 2021; Ikewaki et al., 2022; Thomas et al., 2022) of
their effect on the upper respiratory tract (Zhong Liu, Lu & Xu, 2021),
diabetes (Sivieri, de Oliveira, de Souza Marquez, Pérez-Jiménez, &
Diniz, 2022) and other effects such as antioxidant and antimicrobial
activities (Khan, Date, Chawda, & Patel, 2019), among others. Although
some review works address the inclusion of this by-product in foods
(Ciecierska, Drywien, Hamulka, & Sadkowski, 2019; Rakowska, Sado­
wska, Dybkowska, & Swiderski, 2017; Samuelsen 2014), it was not until
recently that β-glucans isolated from yeasts were reported to be fer­
mented by the gut microbiota (Wang, Chen, Li, Zheng, & Zeng, 2020).
The present review is focused on the biological effect of β-glucans
isolated from brewers spent yeasts in the intestinal context, deepening
the relation of its prebiotic effect and other related biological responses,
such as immunomodulatory and antitumoral. A viewpoint of sustain­
ability and a trend for the reutilization of this by-product coming from a
growing industry in the context of the circular economy will be also
approached.
2. β-glucans in yeast cell walls
Yeasts are unicellular fungi with spherical or elliptic morphology and
they range in length from 3 to 10 μm. The cell wall is multilaminated,
and the lamellae that integrate it are formed by variously oriented fibrils
(Alexopoulos, 1962). Yeasts stand out mainly for their ability to ferment
carbohydrates; hence the name Saccharomicetes (Gr. saccharon = sugar
+ myketes = fungus) which is applied to these organisms. Due to this
property and the resulting alcohol and carbon dioxide, the bakery, wine,
and brewing industries employ yeasts. Cell walls determine the shape of
fungal cells and are essential for their integrity. They consist mainly of
carbohydrates, some free, some linked to proteins.
The β-glucans represent 50–60% of the yeast cell wall composition,
mannoproteins and chitin account for 35–40% and 1–3%, respectively
(Pengkrumsi et al., 2016) and glycogen is around 22% (Deshpande,
Sankh, & Arvindekar, 2011). The β-glucans from S. cerevisiae are
composed of β-(1 → 3) and β-(1 → 6)-linked glucose residues (“bran­
ch-on-branch structure”) (Barsanti et el., 2011), in the proportion of 5:1
(Mantovani et al., 2008).
2
Food Hydrocolloids 133 (2022) 107963
The cell wall of Saccharomyces sp. is described as organized into three
layers (Fig. 1): an external layer containing phosphorylated man­
noproteins, a middle layer containing chains of β-(1 → 6)-glucans and
β-(1 → 3)-glucans, and a rigid inner layer containing chitin (Bastos et al.,
2022; Geoghegan, Steinberg, & Gurr, 2017; Kollár et al., 1997).
The polysaccharides appear to have a structural function, whereas
the mannoproteins may act as “filler” and are important for the
permeability of the yeast cell wall (Kollár et al., 1997). Saccharomyces
cell walls are mostly constituted by β-glucans, with most of the glyco­
sidic linkages β-(1→3) and a small amount of β-(1 → 6). Regarding yeast
glycogen, it occurs in two pools: (1) in the cytosol, on soluble form and
(2) in the cell wall, on insoluble form, covalently linked to β-(1 → 3)
glucans through β-(1 → 3,6) linkages. These linkages are essential to
maintain the components in their right positions, as well as preventing
dissolution/lixiviation of soluble polysaccharides (Deshpande et al.,
2011).
The yeast β-glucans could be classified according to their solubility in
aqueous solutions. In this sense, three (3) groups of β-glucans can be
identified: i) water soluble (β-(1→6) glucans, short polymers); ii) alkali
soluble (β-(1 → 3) glucans not attached to chitin), and iii) alkali insol­
uble (β-(1 → 3, 6) glucans and linked to chitin) which constitutes the
major fraction (Bastos et al., 2022).
As can be concluded, β-glucans on the yeasts cell walls are abundant
but are intimately associated with other biomolecules. Although it is
interesting to use these polymers, care must be taken in the choice of the
isolation method, since this will influence the purity and characteristics
of the β-glucan obtained (Bastos et al., 2022). A discussion regarding
methods of extraction, purity and yield is approached in section 4.
Interestingly, yeasts cell wall is a dynamic structure which vary between
different metabolic stages of the cells, i.e before, during or after the
fermentation (Bastos et al., 2022), which contributes to the complexity
of β-glucans isolation from BSY.
3. Reducing waste: brewers spent yeast: reuse and exploitation
in circular economy
Nowadays there is a political and social great interest to reduce the
pollution arising from industrial activities. Almost all developed and
underdeveloped countries are trying to adapt to this reality by modi­
fying their processes so that their residues can be recycled
Fig. 1. Yeast cell wall structure and distribution of polymers in three (3) layers.
Adapted from Geoghegan et al. (2017).
M.A. Caruso et al. Food Hydrocolloids 133 (2022) 107963

(Jagadiswaran et al., 2021). Consequently, most large companies no

longer consider residues as waste, but as a raw material for other pro­
cesses. The brewing industry generates relatively large amounts of
by-products and wastes such as spent grain, spent hops and yeast being
the most common. However, as most of these are agricultural products
they can be readily reused, as the brewing industry tends to be more
environmentally friendly (Cimini & Moresi, 2021; Silva & Morais,
2021). Indeed, the whole food industry is trying to find new applications
that will change the traditional approach to “waste” products and make
them “co-products” (Farcas et al., 2021). Modern food science and
technology aim to valorize food industry by-products to produce
chemicals, raw materials, and other value-added compounds (Helkar,
Sahoo, & Patil, 2016; Moretti et al., 2022; Rachwał, Waśko, Gustaw, &
Polak-Berecka, 2020).
The spent yeast generated by the brewing industry -compared to
other fermentation industries- is produced in a relatively pure form. It is
used mainly as a feed and a base for seasonings and nutritious agents,
but a considerable part is thrown away. More efficient utilization with
higher added value is therefore desired. Although BSY can be reused to
inoculate subsequent fermentations, this practice is not implemented by
most producers and even doing so, the number of times that yeasts can
be reused is limited (4–16 times); moreover, special knowledge or spe­
cial laboratory conditions are needed to implement this practice safely.
On the other hand, given the volume of yeasts not reused and the dif­
ficulties in disposing them as waste, their final discarding also consti­
tutes an environmental problem that leads to the continuous search for
alternatives. This effluent represents an inconvenience for the brewing
industry, especially due to its high moisture content and its high content
of biodegradable organic matter, which makes it vulnerable to rapid
degradation and thus unstable from the microbiological point of view. Fig. 2. Flowchart of the two main processes conducting the use of BSY: as
When spent yeast is administered to animals, the main destination is whole cells (A) or after cell lysis (B). After cell lysis, two main by-products can
be obtained: intracellular content (so called “yeast extract”, C), and cell wall
as feed for pigs, ruminants, poultry, horses, salmon, and swine (Ferreira
components (D), which contain β glucans.
et al., 2010; Øverland, Karlsson, Mydland, Romarheim, & Skrede, 2013).
Given its high digestibility and its high content of proteins, vitamins
(mainly B group), and minerals (especially phosphorus) it can be used in and/or ultrasound processing. The choice of the method has a consid­
animal diets (Rachwał et al., 2020). Before administration to animals, erable impact on composition and quality of the yeast extract (Jacob,
yeasts must be inactivated by heat treatment or by addition of organic Striegel, Rychlik, Hutzler, & Methner, 2019b; Marson et al., 2020).
acids, in order to avoid gastrointestinal inflammation. Mechanical methods are candidates to be industrially scalable (Jacob
One of the possible applications for the reuse of BSY is the extraction et al., 2019b: Puligundla et al., 2020). Cell lysis methods are
of β-glucans, which are considered pro-health additives. β-Glucans
modulate the immunological response of humans and other animals,
exhibits prebiotic and antioxidant activity, and positively influences
blood lipid content, among others (Lam & Cheung, 2013; Rakowska
et al., 2017). These properties make glucans attractive supplements for
functional foods, and BSY is a good source for the extraction of this
compound. Health properties will be addressed in next sections (5 and
6). It is important to remark that a local or regional governmental pol­
icies of disposal are needed for the implementation of reuse practices in
beer industry (and others) by-products in the circular economy.

4. Methods of extraction

As shown in Fig. 2, BSY could be used as whole cells or can be lysed

by different methods (see below). If lysed, intracellular content (yeast
extract) or cell wall components (mainly polysaccharides) can be used
(Jacob, Striegel, Rychlik, Hutzler, & Methner, 2019a; Watanabe, Inaida,
Yamada, & Karakawa, 1980).
There are several ways to extract the β-glucans from the yeast’s cell
walls. They may differ in terms of the purity of the product obtained and
the environmental cost. The first step is always to disrupt the cell wall.
Cell disruption is sometimes called “autolysis”, despite this is not always
correct since auto means “by the cell”, nevertheless the word is often
used to refer to cell disruption, regardless of how it is done. Anyway, this
procedure has been performed by the so called “mechanical”, or “non-
mechanical” methods, using the induction of cell lysis (pH and high Fig. 3. Classification of usual methods conducted to elicit yeast cell lysis.
temperature-based protocols), glass bead milling, enzymatic hydrolysis Adapted from Puligundla et al. (2020).

3
M.A. Caruso et al. Food Hydrocolloids 133 (2022) 107963

summarized in Fig. 3. Table 1

Cell lysis could be natural or induced; the first one (autolysis) occurs Different available technologies for mechanical Cell Disruption. Comparison of
during alcoholic fermentation and aging process, and it is not used their main characteristics and scalability. Information adapted from Patel,
industrially because it cannot be controlled (Alexandre & Guilloux-Be­ Chakraborty, and Murthy (2016); Yurdacan and Sari (2021) and www.microflui
natier, 2006). Induced cell lysis is wildly used in food and cosmetic in­ dicscorp.com.
dustries, where this technique is used to extract nucleotides, amino acid, Technology Generalities Scalability
and antioxidants from yeast cells (Rakowska et al., 2017). During cell French Press Generates high pressure in a pressured Not
lysis a variety of intracellular materials including large, medium-size, cubicle. A manually controlled valve scalable
and small molecular weight proteins, long chain and short chain fatty releases the pressurized fluid from the
cubicle, resulting in cell rupture.
acids, and polysaccharides are released to the culture medium through
High pressure The most widely used method for both Scalable
passive transportation processes (Alexandre & Guilloux-Benatier, 2006). homogenizers small and large scale due to its high
The addition of exogenous enzymes (mainly proteases) is a usual option, (HPH) efficiency, simplicity and lower cost
as it releases components from the wall in a controlled and efficient compared to other alternatives.
manner, while the process can be scaled-up (Borchani et al., 2016; It involves passing the suspension to be
homogenized through a very narrow
Bzducha-Wróbel et al., 2014).
channel under high pressure.
Regarding mechanical disruption processes, they are carried out in a Ultrasonication Utilizes cavitation forces. An ultrasonic Not
non-specific way, involving, i.e., agitation with glass beads, cavitation probe sonicates the cell suspension. Low scalable
by high pressure or ultrasound and thermolysis (Jacob et al., 2019b). price.
Small sample volumes only. Auditive
The most common method involves glass bead agitation (so called
contamination.
“milling”). Ultrasound techniques are effective but require a large Freeze-thawing Subjecting the cell suspensions to variable Not
amount of energy for long periods of time. Mechanical methods allow to temperatures results in rupture of the scalable
maintain the characteristics of the yeast intracellular components walls. This is not a very reproducible
(including enzymes) stable, and the process can be scaled-up. Table 1 method so results will vary. It is only
suitable for very small samples in the ml
summarizes the advantages and disadvantages of mechanical methods.
range.
Chemical disruption is performed using bases, acids, surfactants, Chemical Lysis Adding chemicals that soften and rupture Limited
detergents, and solvents (Middelberg, 1995; Suwanapong, Khongsay, the cell walls. Chemicals can be expensive
Laopaiboon, Jaisil, & Laopaiboon, 2013). Chemical methods act by and contaminate the preparation which
may be undesirable.
permeabilizing cell wall, which allows intracellular products to pass
Mortar and Pestle Consist on grinding the cell suspension. It Not
through it. These methods have limited potential for scaling up, low is a laborious manual work (not scalable
efficiency and low economic viability; moreover, chemical treatments repeatable) and that can take several
can degrade compounds with biological properties (Liu, Ding, Sun, minutes. Only suitable for small lab
Boussetta, & Vorobiev, 2016). samples
Microfluidizer It uses high fluid pressure to generate Scalable
Following the cell lysis, the intracellular content and cell wall should
shear forces. Achieves a uniform particle
be separated, usually by centrifugation (Pengkumsri et al., 2016; Puli­ size reduction (crystallization,
gundla et al., 2020). Supernatants contain intracellular cell components nanoencapsulation, nano and micro
(“yeast extract”) which have a wide field of applications (for a review, emulsions). Repeatable.
Media Milling Needs special equipment. It tends to be an Limited
see Puligundla et al., 2020), while pellets contain cell walls with
effective way of rupturing many cell types,
β-glucans (reviewed here). Purification is needed since the complete cell but contamination by the media and
wall still contains other components. Purification is done with alkali and temperature control are hazards.
basis treatments (Peltzer, Salvay, Delgado, de la Osa, & Wagner, 2018; Enzyme pre- Enzymes soften the cell walls prior to Limited
Pengkumsri et al., 2016). By this methodology, β-glucans with a high treatment mechanical disruption. This technique can
be valuable when combined with other
percentage of purity could be obtained (Peltzer et al., 2018; Pengkumsri
ones, as it can reduce the pressure or
et al., 2016; Piermaria, Rivero, & Medrano, 2021). According to the number of passes required.
methodology used, extracts of yeast walls with different polymer
composition and different degree of polymerization are obtained.
Therefore, it is convenient to use the appropriate methodology, ac­ and purity, while increasing the cost of the procedure. Therefore, milder
cording to the purpose or the application that the polymers obtained will methods have been developed, avoiding drastic conditions and
have. Otherwise, it is relevant to discriminate the kind of glucans ob­ increasing yield, removing contaminating components (proteins, lipids)
tained after any process of purification, thus avoiding incorrect ex­ as well as mixed methods, which employ hot water, organic solvents and
trapolations to all β-glucans (Bastos et al., 2022). enzymatic treatments, which have been widely used (Freimund, Sauter,
Yeast β-glucans were classified long ago according to their solubility Käppeli, & Dutler, 2003; Sceni, 2021; Wagner et al., 2008). Some of the
in acid or alkaline solutions (Manners & Meyer, 1977), resulting in three used combined methods include: autolysis and alkaline treatment
fractions: 1) alkali-soluble fraction, containing β-(1,3) -D-glucans, (Suphantharika, Khunrae, Thanardkit, & Verduyn, 2003), obtaining a
mannans and some β-(1,6)-D-glucans; 2) acid-insoluble and purity of 51%; or autolysis and high pressure homogenization, plus hot
akali-insoluble fraction, composed of β-(1,3)-D-glucans bound to chitin; acid and alkaline treatment (Thammakiti, Suphantharika & Verduyn,
and 3) alkali-insoluble fraction, constituted by β-(1,6)-D-glucans 2004), obtaining a purity of 59%. Also, thermic, soft alkaline and pro­
(Wagner, Sceni, & Otero Rambla, 2008). On the other hand, in the yeast tease treatments were combined (Freimund et al., 2003) obtaining a
cell wall, the β-(1,3)-D-glucan branched with β-(1,6) inter-chain link, yield of 87% and a purity higher than 92%. Other authors improved
forms the core of the wall, and its non-reducing ends are cross-linked to yield by using a combination of autolysis with mechanical disruption
the reducing end of chitin chains through a β-(1,4) linkage, and this such as high pressure homogenization (Liu, Wang, Cui, & Liu, 2008) or
complex is bound to β-(1,6)-D-glucans or mannoproteins. Such structure ultrasound treatment (Da Silva et al., 2014; Magnani et al., 2009) fol­
makes the polysaccharide insoluble in both water and hot alkali lowed by organic solvents extraction, obtaining not-degraded polymers
(Aimanianda et al., 2009; Barsanti et al., 2011; Sugawara, Takahashi, with a purity higher than 93%. High pressure homogenization followed
Osumi, & Ohno, 2004). by autolysis also allowed an acceptable yield (Dimopoulos, Tsantes, &
The use of the above mentioned extraction methods results in Taoukis, 2020), while the combination of thermal and enzymatic
different insoluble residues, which can be degraded, thus reducing yield treatments resulted in a decrease of yield, but acceptable purity;

4
M.A. Caruso et al. Food Hydrocolloids 133 (2022) 107963

enzymatic treatments are not preferred when scalability is projected, Table 2

due to the high cost and the lack of recovery of the enzymes (Borchani Biological activity of β-glucans isolated from BSY, assorted by type of biological
et al., 2014), despite them contribute to the solubilisation of the polymer activity and then by model (in vitro vs in vivo and trials, if done).
(Pinto, Coelho, Nunes, Brandão, & Coimbra, 2015). Abbreviations: GIT: gastrointestinal tract; SCFA: short chain fatty acids. mAbs:
It is concluded that the type of extraction modifies the physical and monoclonal antibodies.
structural properties of the obtained glucans, and therefore, the bio­ 1 - PREBIOTIC
logical activity of the polymer is affected (Bastos et al., 2022; Chaiyasut Model Main Results Reference 1
et al., 2018; Hromádková et al., 2003; Pizarro, Ronco, & Gotteland,
Human faecal slurries Incubation of faecal slurries Chaikliang et al. (2015)
2014). There are still no taxonomic studies that correlate methods and in vitro with β-glucans increased the
products, maybe because more results are required to reach conclusions Prebiotic Index. Acetate was
on this regard. On this sense, unfortunately, there is not enough scien­ the most prevalent SCFA found,
tific information which allows to connect extraction protocols and mo­ followed by propionate,
butyrate and lactate.
lecular structure (degree of branching, ratio β-1-3/1–4 linkages, GIT passage β-glucan was not degradable by Wang et al. (2020)
molecular weight, conformational effects, solubility, degree of purity) simulation - human GIT enzymes and stimulated
with biological properties of the obtained polymer, although it should be faecal slurries in beneficial bacteria populations
relevant (Bastos et al., 2022; Zhu, Du, & Xu, 2016). vitro such as Bifidobacterium longum,
while inhibited the
proliferation of harmful gut
5. β-glucans from S. cerevisiae: biological activity microbiota.
Fish (Solea Orally administrated insoluble Carballo et al. (2019)
The biological activity of fungal β-glucans in general is well recog­ senegalensis) in vivo yeast β-glucan acted locally in
the gut of the fishes, controlling
nized and scientific reports on their beneficial properties for health are
Vibrio genus abundance and
increasing (Cerletti et al., 2021; Murphy et al., 2020; Nakashima et al., enhancing beneficial bacteria.
2018; Thomas et al., 2022). Fungal β-glucans are considered prebiotics Piglets (administered Oral supplementation of yeast De Vries et al. (2020)
which stimulate the growth and activity of healthy intestinal microbiota, from day two after β-glucans conducted to modest
while inhibiting the growth of pathogens, preventing inflammation as birth until two effects on faecal microbiota
weeks after composition regarding
well as colon cancer (Khan, Date, Chawda & Pate, 2019). Recently, it has
weaning) in vivo diversity indexes. The genus
been shown that β-glucans obtained from fungal cell walls cross the Romboustia was increased and
digestive system without being hydrolysed and that they can be fer­ correlated with dendritic cells
mented by the intestinal microbiota, favouring the development of maturation markers.
2 IMMUNOMODULATING
beneficial bacteria such as Bifidobacterium longum, which is why they are
Shrimps (Penaeus In vitro, β-glucan significantly Suphantharika et al.
considered prebiotic compounds comparable to inulin and have the monodon) enhanced phenoloxidase (PO) (2003)
potential to be considered functional food ingredients (Wang, Chen, Li, in vitro and in vivo activity of shrimps hemolymph.
Zheng, & Zeng, 2020). In vivo, oral administration of
Moreover, fungal β-glucans are frequently called as “nutraceuticals” 0.2% (w/w) of β-glucan for 3
days increased the PO-activity
(Bacha, Nasir, Iqbal, & Anjum, 2017; Ciecierska et al., 2019; Rakowska
of the shrimp.
et al., 2017) since they are a pharmaceutical alternative which claims Zebrafish embryonic In vitro and in vivo: increment Medina-Gali et al.
physiological benefits. In the US, “nutraceuticals” are largely unregu­ fibroblast ZF4 cells on the survival and viability of (2018)
lated, as they exist in the same category as dietary supplements and food and zebrafishs in cells against SVCV (spring
additives by the FDA (Food and Drug Administration), under the authority vitro and in vivo viremia of carp virus)
In vivo: changes on gene
of the Federal Food, Drug, and Cosmetic Act (Web Reference 2). β-glu­ expression of innate immunity
cans were also recognized as safe food additives by the European Food cells.
Safety Authority (EFSA Panel on Dietetic Products, 2011). Moreover, Fishes (Solea Orally administered Carballo et al. (2019)
β-glucans were also been called as Biological Response Modifiers (BRM), senegalensis) in vivo commercial β-glucans
modulated the innate immune
since mainly activate immune system cells such as macrophages (Puli­
response in Senegalese sole
gundla et al., 2020). (Solea senegalensis)
β-glucans from yeasts provide multi-directional biological effects. Pacu (Piaractus β-glucans modulated cortisol Lopes et al. (2022)
(1–3), (1–6)-β-D-glucans from S. cerevisiae are well-known immuno­ mesopotamicus) profile, reducing the levels
modulators in human and animal models, with anti-inflammatory ef­ in vivo after inoculation with
Aeromonas hydrophila,
fects, and were studied as adjuvants in conventional chemotherapy confirming its
(Barsanti et al., 2011; Bastos et al., 2022; Fuller et al., 2017; Murphy imunostimulating action, by
et al., 2020; Wang et al., 2020). Moreover, there is an increasing interest increasing the respiratory
in the recent years for novel pharmaceutical and biomedical applica­ activity of leukocytes and
lysozyme levels after 24 h of
tions, with an emerging focus in immunology, tissue engineering, vac­
inoculation.
cines, and drug delivery systems (Bastos et al., 2022). Yeast derived Broiler chickens Dietary β-glucan Omara et al. (2021)
β-glucans have been approved by the European Food Safety Authority in vivo supplementation to chicken
(EFSA Panel on Dietetic Products 2011) and US Food and Drug diets modulated their immune
Administration (Web Reference 3) as safe novel food ingredients, and response to the Eimeria
challenge (coccidiosis)
β-glucans are already being commercialized as dietary supplements in evaluated on IL10, IFNɣ and
more than 30 countries (Web Reference 4). The patented ingredient has macrophage migration.
broad regulatory approval, including GRAS (Generally Recognized As Broiler chikens Yeast β-glucans Tian et al. (2016)
Safe) from the US Food and Drug Administration and China (Web in vivo supplementation decreased
Clostridium perfringens-induced
Reference 5). The biological activities of yeast β-glucans are summarized
necrotic enteritis lesions and
in Table 2 and will be sddressed in the following paragraphs. As can be C. perfringens colonization.
noted, applications of yeast β-glucans as Biological Response Modifiers Yeast β-glucans
include not only humans and other mammals (mice, pigs), but also supplementation enhanced
include applications in poultry, aquiculture and invertebrates, giving (continued on next page)

5
M.A. Caruso et al.
Table 2 (continued )
1 - PREBIOTIC
Model Main Results
humoral immunity and
improved intestinal health.
Mice β-D-glucan intragastrical
S180-bearing mice administrated potentiated the
(Kunming SPF male mouse immune responses by
mice) decreasing the ratio of CD4 to
in vivo CD8 and increasing the
expression levels of IL-2, IL-6
and TNF-α.
β-D-glucan enhances the host’s
immune function during the
tumour inhibition process.
Mice (C3H/Hej mice) Intraperitoneal injection
in vivo increased immunological
activity as seen in increased
natural killer (NK) and
lymphokine-activated killer
(LAK) activities
Mice (macrophages β-D-glucan increased TNF-α
from mice of ICR induction activity in the murine
line) ex vivo macrophage model.
Piglets (administered Oral administration of
from day two after β-glucans did not modified
birth until two vaccine response in piglets, and
weeks after a moderate immune response
weaning) was observed. The increased
in vivo production of the anti-
inflammatory cytokine IL-10 by
MLN cells and the decreased
production of TNFα by PBMCs,
suggests an anti-inflammatory
effect.
Human clinical trials Orally administrated yeast
β-glucans had positive effects
on upper respiratory tract
infections.
3 ANTITUMORAL (*)
Human monocytes Macrophages treated with
differentiated into curdlan, yeast-b or zymosan
macrophages demonstrated enhanced
coincubated with production of chemo-
patient melanoma attractants, such as CCL3,
tissues CCL4, and CXCL8, which
in vitro contribute to recruitment of
monocytes and neutrophils.
The secretion of chemo-
attractants was confirmed
when using patient-derived
melanoma-infiltrating immune
cells. Taken together, the
bacterial-derived curdlan as
well as the yeast-derived
β-glucans yeast-b and zymosan
have the unique ability to
preferentially skew
macrophages towards a chemo-
attractant-producing
phenotype that may aid in anti-
cancer immune responses.
cell line: Huh7; mice: Liver cancer cells: inhibits
C57BL/6 and autophagy and lysosomal
BALB/c function of cancer cells; inhibits
in vitro and in vivo cell proliferation and
metabolism, and conducting to
apoptosis in hepatocellular
carcinoma.
Mice liver: Inhibits tumour
growth.
Mice Intravenous administration of
BALB/c or C57Bl/6 β-glucan conducted to adjuvant
Induced tumors in activity for antitumor mAbs,
vivo recruitment of granulocytes to
tumor microenvionment and
reduction of tumour growth.
Table 2 (continued )
Reference 1
Mo et al. (2017)
Gu et al. (2005)
Liepins et al. (2015)
De Vries et al. (2020)
Fuller et al., 2017;
Murphy et al., 2020;
Zhong et al., 2021
De Graaf et al. (2021)
Wang, Liu, et al. (2020)
Hong et al. (2003)
6
Food Hydrocolloids 133 (2022) 107963
1 - PREBIOTIC
Model Main Results Reference 1
Mice Orally administered particulate Li et al. (2010)
in vivo β-glucan trafficked into spleen
and lymph nodes and activated
DCs that captured dying tumor
cells in vivo, leading to the
expansion and activation of
antigen-specific CD4 and CD8 T
cells. In addition, IFN-γ
production of tumor-
infiltrating T cells and CTL
responses were significantly
enhanced on β-glucan
treatment, which ultimately
resulted in significantly
reduced tumor burden.
Mice (C3H/Hej mice) Intraperitoneal injection of Gu et al. (2005)
in vivo β-glucan induces a delay on
tumour growth. Natural killer
(NK) and lymphokine-activated
killer (LAK) cells activities were
significantly increased by
repeated doses of β-glucan.
Mice (S180-bearing β-D-glucan intragastrical Mo et al. (2017)
mice administration conducted to:
Kunming SPF male → Decreased tumor volume
mice) in vivo → Increase in the proportion of
apoptotic cells
4 OTHER BIOLOGICAL EFFECTS
Antioxidant
In vitro The antioxidative activity of Jaehrig et al. (2008)
glucan from the cell walls of
S. cerevisiae grown on different
media (wort,
yeast–peptone–glucose,
yeast–peptone–galactose) was
investigated. The results show
significant differences in the
glucan content of the cell walls
and in their antioxidative
activities depending on growth
medium. However, glucan
itself seems to have a low
antioxidative activity in
contrast to other cell wall
fractions e.g. proteins.
Rats (Lewis type male Carboximetilated beer yeast Kogan et al. (2005)
rats) β-glucan showed antioxidant
Experimental activity in the arthritis model,
model of induced suggesting possible application
arthritis of the yeast glucan derivatives
in vivo in the treatment of arthritis.
Antibacterial and Toxin complexes
in vitro Cell walls containing high Yiannikouris et al.
percent of β-glucans were able (2004)
to complex with mycotoxins
(Zearalenone) and thus limiting
their bioavailability in animal
feed when passing the GIT
in vitro The antibacterial activity of Khan et al. (2016)
modified yeast β-D-glucan was
evaluated using disc diffusion
method against 2 g positive
bacteria (Bacillus cereus and
Staphylococcus aureus) and 3 g
negative bacteria (Escherichia
coli, Salmonella typhimurium &
Proteus vulgaris).
Hypocholesterololemic
Rats The diets supplemented with Waszkiewicz-Robak
Wistar both additives were more and Bartnikowska
in vivo effective in lowering (2009)
concentration of serum total
cholesterol as well as LDL-
cholesterol and triacylglycerols
(continued on next page)
M.A. Caruso et al. Food Hydrocolloids 133 (2022) 107963

Table 2 (continued ) metabolized by the microorganisms inhabiting in the large intestine,

1 - PREBIOTIC exerting a regulatory effect, promoting the development of beneficial
microbiota and inhibiting the proliferation of harmful bacteria (Tamura
Model Main Results Reference 1
et al., 2017; Wang, Chen, et al., 2020). Moreover, β-glucans were more
in experimental rats than the able to promote the growth of B. longum compared to inulin. Therefore,
control diet. In conclusion,
β-glucans showed a similar prebiotic activity to inulin and is expected to
feeding rats with dried, spent
brewer’s yeast or β-glucans be developed as a functional food. Nevertheless, an in situ comparison
may significantly reduce about the effect of both polymers into the intestinal context is still
cholesterol accumulation in needed.
liver. Positive effect on beneficial microbiota is one of the targets for new
Porcine model Coronary arteries protection Aarsæther et al. (2012)
in vivo Oral β-glucan pre-treatment
studies for the explanation of healthy effect. Indeed, scientific knowl­
reduces infarction size and edge about prebiotic effect of BSY β-glucans are increasing (Carballo
improves regional contractile et al., 2019; Chaikliang, Wichienchot, Youravoug, & Potchanapond,
function in a porcine 2015; da Silva Guedes et al., 2019; De Vries et al., 2020; Wang, Chen,
ischaemia/reperfusion model.
et al., 2020). In this context, it is important to correlate the specific
Mice (C57BL/6 mice) Oral supplement formulation, Abou Nehmi et al.
in vivo containing prebiotics, yeast (2021) groups of the microbial communities associated with β-glucans
β-glucans, minerals and fermentation and the consequential Short Chain Fatty Acids (SCFA)
silymarin (Silybum marianum), profiles. In this sense, commercial β-glucans from S. cerevisiae (as other
improved lipid and glycidic β-glucans coming from edible mushrooms) induced the production of
metabolism, inflammatory and
mitochondrial proteins of the
acetate as the most prevalent SCFA followed by propionate, butyrate
liver, in control and high-fat and lactate by microorganisms present in human faecal slurries
diet-induced obese mice. (Medrano, Simonelli, Miller, Piermaria & Abraham, 2020; Chairlikang
Pacu (Piaractus β-glucan showed a modulatory Lopes et al. (2022) et al., 2015). This SCFA relative abundance is in concordance to other
mesopotamicus) in effect on lipid metabolism in
prebiotic polysaccharides studied in equivalent in vitro models
vivo fish. We found a reduction in
plasma triglyceride levels, as (Medrano, Gangoiti, Simonelli, & Abraham, 2020). Through the stim­
also in lipid levels in liver, ulation of healthy microbiota and bioactive SCFA, a systemic healthy
muscle, and visceral depots, in effect could be expected to be elicited by these polymers. There are not
fish fed with β-glucan. The yet studies related to SCFA profiles obtained after in vitro faecal slurries
triglyceride levels decreased by
32.9% in fish fed with 0.1%
fermentation and type or structure of β-glucans.
β-glucan, when compared to
the control group. Lipid levels 5.2. Immunomodulating
decreased by 42.8% and 26.8%
in the liver, and by 37.9% and
One of the most studied biological activities of yeast β-glucans is
24.1% in muscle, for fish fed
with 0.1% and 0.5% β-glucan, their immunomodulating property (Ciecierska et al., 2019; Samuelsen,
respectively, compared to the Schrezenmeir, & Knutsen, 2014), which has been addressed in different
control group. Visceral fat models on in vitro and in vivo studies, and even in clinical trials, as shown
decreased by 32.1% and
in Table 2. In vivo studies were performed in Arthropoda (crustaceans)
46.4%, in fish fed with 0.1%
and 0.5% β-glucan,
such as shrimps (Suphantharika et al., 2003; Thanardkit, Khunrae,
respectively. Suphantharika, & Verduyn, 2002), in vertebrates including fishes such
Wound healing as: zebra fish (Medina-Gali et al., 2018), senegalese sole (Solea senegal­
Human trial The effects of the glucan on Medeiros et al. (2012) ensis) (Carballo et al., 2019), or pacu (Piaractus mesopotamicus) (Lopes,
wound healing were assessed in
de Mello, & Urbinati, 2022), and other vertebrates as broiler chickens
human venous ulcers by
histopathological analysis after (Omara, Pender, White, & Dalloul, 2021; Tian, Shao, Wang, & Guo,
30 days of topical treatment. (1 2016), pigs (De Vries et al., 2020) and mice (Chaiyasut et al., 2018;
→ 3)-β-glucan enhanced ulcer Hofer & Pospísi, 1997; Li et al., 2010; Liepins et al., 2015). The main
healing and increased
results of immunomodulatory effects found in these experimental
epithelial hyperplasia, as well
as increased inflammatory
models are summarized in Table 2. Clinical trials indicated that the
cells, angiogenesis and orally administrated yeast derived β-glucans had positive effects on
fibroblast proliferation. upper respiratory tract infections (Fuller et al., 2017; Murphy et al.,
(*) When antitumoral studies were related to immunomodulatory effect (and in 2020; Zhong, Liu, Lu, & Xu, 2021). Moreover, a clinical trial is currently
order to not repeat the information), the bibliographic reference was cited only on-going related to the effect of β-glucan-containing supplement (called
once, taking into account the biological effect prioritized by authors in their ABBC1) on volunteers receiving the Influenza or Covid-19 vaccine
original research. (Geller & Yan, 2020; Xavier-Santos et al., 2022).
β-glucans seems to elicit a positive effect on immune function,
account for the wide range of applications that theese polymers have in nevertheless the mechanisms of action remain still unknown. It was
different animal models, thus expanding the uses that can be given to proposed that immunomodulation could be related to the ability to
this by-product. It is worth to be said that the structural variability of modulate the gut microbiome (Carballo et al., 2019; De Vries et al.,
yeast β-glucans (molecular weight, type of branches and degree, solu­ 2020). Different studies postulate than the whole molecule can trigger
bility, purity among others) affect their biological activity (Bastos et al., the biological effects by itself. It was hypothesized that the interaction of
2022). the molecule with mammalian cell receptors such as Dectin-1 and
Complement Receptor type 3 (CR3) (both expressed in immune cells)
are responsible of triggering the immunomodulatory event, by binding
5.1. Prebiotic and activation (De Marco Castro, Calder & Roche, 2021; Pan et al.,
2019). This could be relevant by the application of β-glucans as adju­
β-glucans could pass through the digestive system without being vants of immune responses (Bastos et al., 2022). Phagocytosis by mac­
hydrolysed by mammalian digestive enzymes. β-glucans could then be rophages was also reported as a mechanism of action of β-glucans in

7
M.A. Caruso et al.
mice and humans, which allows to elicit a systemic response (Li et al.,
2010; Pan et al., 2019). As insoluble β-glucans seem to be “more
bioactive” than soluble ones -at least regarding immunomodulatory
properties as adjuvant-chemical modifications have been applied to
improve such biological property (Bastos et al., 2022). Both the effect
through the organic acids produced by the microbiota when the polymer
is orally administered, as well as the stimulation of the immune system
when the polymer is used as an adjuvant, are promising applications of
β-glucans as stimulants of the immune system and as modifiers of the
biological response.
5.3. Antitumoral
Through stimulation of the immune system by itself or/and stimu­
lation of gut microbiota, yeast β-glucans have been shown to have
antitumoral effect (Ciecierska et al., 2019; Mantovani et al., 2008;
Vetvicka & Vetvickova, 2018), which was demonstrated in mice and
rats’ models (Gu et at., 2005; Hong et al., 2003; Mo et al., 2017; Wang,
Liu, et al., 2020) or even for humans (Ikewaki et al., 2022; Murphy et al.,
2020). As shown in Table 2, in the case of animal studies, the common
denominator seems to be that the size of tumors is reduced (Gu et al.,
2005; Hong et al., 2003; Mo et al., 2017; Wang, Liu, et al., 2020), and
although the mechanism of action is not entirely elucidated, it could be
proposed that the stimulation of TNFα release is implicated (Liepins
et al., 2015) since β-glucans binds and activates NK cells (Hong et al.,
2003; Petravić-Tominac et al., 2010), which would influence tumour
reduction. Stimulation of macrophages after phagocytosis has been
purposed as a possible mechanism of action, together with the later
stimulation of the adaptative immune system (Li et al., 2006, 2010; Pan
et al., 2019). Modulation of macrophage populations (M1 and M2) was
also purposed as mechanism of action of β-glucans in tumour microen­
vironment (de Graaff et al., 2021). An alternative explanation could be
the production of metabolites with antitumoral activity such as butyrate,
when the polymer is orally administered and is fermented by colon
microbiota (Chairlikang et al., 2015; Medrano et al., 2020b). Addi­
tionally, a direct antineoplasic effect in situ elicited by the polymer
among tumoral cells should not be discarded, i.e: neoplasic colonocytes
(Caruso, Abraham, & Medrano, 2021, November), as it was purposed by
other fungal β-glucans (Khan et al., 2019), even though more in vitro and
in vivo studies are still needed.
5.4. Other Healthy effects
β-glucans isolated from SBY were found to be effective at lowering
the levels of LDL cholesterol and triacylglycerols as well as serum total
cholesterol in experimental mice and rats (Abou Nehmi et al., 2021;
Waszkiewicz-Robak & Bartnikowska, 2009), and protecting the cardiac
system in coronary artery bypass in pigs (Aarsæther, Straumbotn,
Rösner, & Busund, 2012). Thus, β-glucans were proposed as useful di­
etary supplements in diabetes mellitus (Sivieri et al., 2022). Antioxidant
effect was also reported in vitro (Jaehrig et al., 2008; Kogan et al., 2005),
as well as antibacterial activity against Bacillus cereus, Staphylococcus
aureus, Escherichia coli, Salmonella typhimurium and Proteus vulgaris
(Khan et al., 2016), while adsorption of mycotoxins has also been
demonstrated, with the β-glucans acting as binders (Hamza et al., 2019;
Yiannikouris et al., 2004).
Accompanying the aforementioned immunomodulatory activity,
anti-inflammatory properties are related and together with them, the
opportunity to use β-glucans in applications related to tissue engineering
such as bone implants and tissue repair has been suggested (Bastos et al.,
2022). Moreover, wound healing is another promising reported bio­
logical effect (Du, Bian, & Xu, 2014; Majtan & Jesenak, 2018; Medeiros
et al., 2012; Vetvicka & Vetvickova, 2011). This property, together with
new insights into the importance of favouring the skin microbiome,
make these polymers candidates for the development of new ingredients
for the cosmetics industry (Natakankitkul, 2016).
8
Food Hydrocolloids 133 (2022) 107963
It should be concluded that some of the aforementioned health-
related effects of β-glucans could be attributed to the property of
reaching the large intestine where they can act biologically in a direct or
indirect manner, as proposed for other polysaccharides (Sharma, Kata­
ria, Sharma, & Singh, 2022; Simonelli, Gagliarini, Medrano, Piermaria,
& Abraham, 2020; Teferra, 2021). The beneficial effects of the oral
consumption of β-glucans on health are related to their property of
reaching the large intestine and they may be functioning locally due to
the interaction of the polymer with the cells of the intestinal epithelium
and/or with the resident microbiota or systemic, through the metabo­
lites of bacterial populations and/or their interaction with cells of the
immune system (Fig. 4).
Focusing on the underlying mechanisms by which yeast β-glucans
could elicit a biological effect on the intestinal context, some hypothesis
could be done. First, it was demonstrated that the polymer reaches the
colon (Wang, Chen, et al., 2020). Despite the direct interaction of
β-glucans with intestinal epithelial cells was not yet investigated, it
could be expected to occur, since the interaction with mammalian cell
receptors (Dectin-1 and CR3) was demonstrated (De Marco Castro et al.,
2021; Pan et al., 2019). Hydrophobicity of the polymer could be
involved in the close interaction with the cell surface, as was docu­
mented for other polymers (Ayala et al., 2011). In situ, β-glucans could
be fermented by beneficial intestinal microbiota, as demonstrated
(Carballo et al., 2019; Chaikliang et al., 2015; da Silva Guedes et al.,
2019; De Vries et al., 2020; Wang, Chen, et al., 2020). Proliferation of
beneficial microorganisms could inhibit the occurrence of
non-beneficial ones (by competitive exclusion and/or by the acidifica­
tion of the microenvironment), and this could explain the observed ef­
fect against intestinal pathogens (Khan et al., 2016). Direct interaction
of the polymer with the bacterial cell surfaces or their toxins should not
be discarded, since adsorption of mycotoxins has been demonstrated
(Hamza et al., 2019). Additionally, SCFA -product of intestinal micro­
biota fermentation- could be produced and could have a positive effect
in situ (i.e: antiproliferative effect elicited by butyric acid) or by a sys­
temic effect when the SCFA reache systemic blood via the portal vein.
6. Application in human foods - legislation
The legal options for using yeast-derived β-glucans as a novel food
ingredient come under the Regulation of the European Parliament and
Council (EU) 2015/2283 from November 25, 2015 concerning novel
foods, amending the Regulation of the European Parliament and of the
Council (EU) No 1169/2011 and repealing Regulation (EC) No 258/97
of the European Parliament and Council along with Commission Regu­
lation (EC) No 1852/2001 (Rakowska et al., 2017). In the United States
they are included into the same category as dietary supplements and
food additives by the FDA (Food and Drug Administration), under the
legislation of the Federal Food, Drug, and Cosmetic Act (Web Reference
2). In Argentina, there is a precedent for the inclusion of the first beer
by-product (bagasse) in the Argentinian Codex Alimentarius CAA (Web
Reference 6).
Yeast β-glucans are commercialized in more than 20 countries as
dietary supplements to be used in fruit drinks, cereal bars, biscuits,
crackers, breakfast cereals, yoghurt, chocolate, soups, protein bars and
foodstuffs intended for nutritional uses; excepting infant formulae
(Rakowska et al., 2017). Introducing β-glucan into foodstuffs bestows
upon them the characteristics of functional foods.
According to the literature, β-glucans with molecular weight from 16
kDa are insoluble polymers (Mantovani et al., 2008). Extraction and
purification conditions affect the physicochemical properties of glucans.
Then, the methodology used to obtain them must be chosen according to
the application desired to give to the polymer. It was found that ho­
mogenization of brewer’s yeast cell walls improved β-glucan content
and apparent viscosity of the products, while spray drying adversely
affected the apparent viscosity of the β-glucan products. β-glucans ob­
tained from BSY exhibited higher apparent viscosity, water-holding
M.A. Caruso et al. Food Hydrocolloids 133 (2022) 107963

Fig. 4. Purposed mechanisms of action of β -glucans related to their biological activity. Two (2) non-exclusive main interaction pathways are purposed: the direct
interaction of the polymer with host cells, and interaction of the polymer with host microbiome. At the same time, depending on the kind of interaction, response
could be local or systemic. Abbreviations: SCFA: short chain fatty acids: GIT: gastrointestinal tract. Template downloaded from www.Canva.com.

capacity and emulsion stabilizing capacity than commercial purified

Table 3
β-glucans, but similar oil-binding capacity (Thammakiti, Suphantharika,
Functionality of β-glucans in human foods and formulations.
Phaesuwan, & Verduyn, 2004).
Brewer’s yeast β-glucans show a great potential for use as a thick­ Function Food containing β-glucans Reference

ener, water-holding agent, oil-binding agent or emulsifying stabilizer Dietary supplement Allowed for: fruit drinks, cereal Rakowska et al.,
and as a fat replacer in food products such as soups, sauces, desserts and bars, biscuits, crackers, (2017)
breakfast cereals, yoghurt, Samuelsen et al.,
salad dressings (Rachwał et al., 2020; Thammakiti et al., 2004); more­
chocolate, soups and protein (2014)
over, they are useful as wall materials to encapsulate compounds with bars (except for infant formula). Zhu et al., 2016
functional or biological properties (Salgado, Rodríguez-Rojo, & Cocero, According to Regulation of the
2017). European Parliament and of the
Application of yeast β-glucans in food products and formulations are Council (EU) No
1169/2011 and repealing
summarized in Table 3 β-glucans have been used in yoghurts (Pio­
Regulation (EC) No 258/97 of
trowska, Waszkiewicz-Robak, & Świderski, 2009; Raikos, Grant, Hayes, the European Parliament and
& Ranawana, 2018), as functional ingredient in bread (Kittisuban, Rit­ Council along with Commission
thiruangdej, & Suphantharika, 2014; Martins et al., 2015; Rakowska Regulation (EC) No 1852/2001
Thickener, water-holding Soups, sauces Rakowska et al.,
et al., 2017; Rieder, Ballance, Böcker, & Knutsen, 2018), and as a fat
agent (2017)
replacer in mayonnaise formulations (Marinescu, Stoicescu, & Patrascu, Incremental effect on the Yoghurts Piotrowska et al.,
2011; Worrasinchai, Suphantharika, Pinjai, & Jamnong, 2006). More­ textural properties (2009)
over, these polysaccharides can be used as low-calorie raw materials to Raikos et al.,
replace high-calorie raw materials in foods to control or prevent obesity (2018)
Functional ingredient Bread Martins et al.,
and diabetes (Yoo & Lee, 2007). The film-forming property is another
(2015)
interesting physicochemical aspect of this polymer which broadens the Rieder et al.,
field of its applications in food industry (Novák et al., 2012; Peltzer (2018)
et al., 2018). Finally, yeast cell walls in microcapsules, essentially con­ Kittisuban et al.,
(2014)
sisting of 80% β-glucans, have been used for drug encapsulation,
Oil-binding agent or Desserts and salad dressings Thammakiti
showing diverse applications such as delivery of drugs, nanoparticles, emulsifying stabilizer et al., (2004)
vaccines, and diagnostic agents, among others (Bastos et al., 2022; Sabu, and as a fat replacer
Rejo, Kotta, & Pramod, 2018). Fat replacer Mayonnaise Marinescu et al.,
However, as said, these capacities of β-glucans were shown to be (2011)
Worrasinchai
affected by differences in isolation and drying procedures employed for
et al., (2006)
the β-glucan preparation (Bastos et al., 2022; Puligundla et al., 2020). Film forming – Novák et al.,
With the increasing perfection of laws and regulations and the demand 2012
of consumers and markets, yeast cell wall polysaccharides will have Peltzer et al.,
(2018)
more applications in dairy and functional foods, especially the
Encapsulating agent – Sabu et al.,
middle-aged and old formula milk powder, functional fermented yogurt, (2018)
meal powder, biscuits, etc (Rachwał et al., 2020; Zhu, Du & Xu, 2016).

9
M.A. Caruso et al.
7. Conclusions
Craft beer brewery is a growing economic activity in many countries.
One of the most important by-products generated by this economic ac­
tivity is BSY. As demonstrated, BSY and β-glucans isolated from yeast
cell walls have many potential applications to be used as food additives
with positive impact on human heath such as prebiotic, immunomod­
ulatory, antitumoral, anti-inflammatory, and antioxidant functions,
among others. Besides, β-glucans extracted from BSY have interesting
functional properties to be used as food additives. As a consequence of
the increasing interest in circular economy by the whole food industry
(with the aim to be more environmentally friendly by reducing waste
and finding new added value for the generated by-products), there is a
need to identify the means to valorize such by-products and explore
their potential applications as functional ingredients. Scaling up pro­
cesses are desired to be set-up and actualization of the Codex Ali­
mentarius of the involved countries are also needed. The sustainable
production is a new challenging route the scientists and brewers have to
develop.
Authors contributions
Agustina Caruso: Investigation, Writing original draft, Data cura­
tion, Writing - review & editing.
Judith A. Piermaria: Investigation; Validation; Writing - review &
editing.
Micaela Medrano: Conceptualization; Investigation; Supervision;
Resources; Writing - review & editing.
Analía G. Abraham: Supervision; Resources; Writing - review &
editing.
Funding source
This work was supported by the ANPCyT (Grant Number PICT-2016-
0639); CONICET, National University Arturo Jauretche (UNAJ) and
National University of La Plata (UNLP) (Grant Number PIO 2021-2023
044 and UNAJ Investiga 2020).
Declaration of competing interest
None.
Acknowledgments
Agustina Caruso is a fellow of CIN (Consejo Interuniversitario
Nacional). Judith Piermaria, Analía Abraham and Micaela Medrano are
members of the Researcher Career of CONICET (Consejo Nacional de
Investigaciones Científicas y Tecnológicas). Authors would like to
acknowledge to craft breweries of the region to their contribution with
BSY. Authors also want to acknowledge to Ing. Martin Ducos (IPATEC)
for the advisement on the relevance of the subject and for sharing Data
Bases to elaborate surveys to the craft breweries of the region.
References
Aarsæther, E., Straumbotn, E., Rösner, A., & Busund, R. (2012). Oral β-glucan reduces
infarction size and improves regional contractile function in a porcine ischaemia/
reperfusion model. European Journal of Cardio-Thoracic Surgery, 41(4), 919–925.
Abou Nehmi, V., Murata, G. M., de Moraes, R. C. M., Lima, G. C. A., De Miranda, D. A.,
Radloff, K., et al. (2021). A novel supplement with yeast β-glucan, prebiotic,
minerals and Silybum marianum synergistically modulates metabolic and
inflammatory pathways and improves steatosis in obese mice. Journal of integrative
medicine, 19(5), 439–450.
Aimanianda, V., Clavaud, C., Simenel, C., Fontaine, T., Delepierre, M., & Latgé, J. P.
(2009). Cell wall β-(1, 6)-glucan of Saccharomyces cerevisiae: Structural
characterization and in situ synthesis. Journal of Biological Chemistry, 284(20),
13401–13412.
Alexandre, H., & Guilloux-Benatier, M. (2006). Yeast autolysis in sparkling wine–a
review. Australian Journal of Grape and Wine Research, 12(2), 119–127.
10
Food Hydrocolloids 133 (2022) 107963
Alexopoulos, C. J. (1962). Introductory mycology (2nd ed.). London: John Wiley and Sons
(Chapter 6).
Alsteens, D., Dupres, V., Mc Evoy, K., Wildling, L., Gruber, H. J., & Dufrêne, Y. F. (2008).
Structure, cell wall elasticity and polysaccharide properties of living yeast cells, as
probed by AFM. Nanotechnology, 19(38). Article 384005.
Ayala, R., Zhang, C., Yang, D., Hwang, Y., Aung, A., Shroff, S. S., et al. (2011).
Engineering the cell–material interface for controlling stem cell adhesion, migration,
and differentiation. Biomaterials, 32(15), 3700–3711.
Bacha, U., Nasir, M., Iqbal, S., & Anjum, A. A. (2017). Nutraceutical, anti-inflammatory,
and immune modulatory effects of β-glucan isolated from yeast. BioMed Research
International, 2017.
Barsanti, L., Passarelli, V., Evangelista, V., Frassanito, A. M., & Gualtieri, P. (2011).
Chemistry, physico-chemistry and applications linked to biological activities of
β-glucans. Natural Product Reports, 28(3), 457–466.
Bastos, R., Oliveira, P. G., Gaspar, V. M., Mano, J. F., Coimbra, M. A., & Coelho, E.
(2022). Brewer’s yeast polysaccharides—a review of their exquisite structural
features and biomedical applications. Carbohydrate Polymers, 277. Article 118826.
Boateng, M., Okai, D. B., Frimpong, Y. O., & Zeebone, Y. Y. (2015). Wet brewers’ spent
grains and wet brewers’ spent yeast: Problems associated with their usage and
suggested solutions: A case study of the Ejisu-Juaben Municipality of Ghana.
Livestock Research for Rural Development, 27. Article 5.
Borchani, C., Fonteyn, F., Jamin, G., Paquot, M., Blecker, C., & Thonart, P. (2014).
Enzymatic process for the fractionation of baker’s yeast cell wall (Saccharomyces
cerevisiae). Elsevier Food Chemistry, 163, 108–1.
Borchani, C., Fonteyn, F., Jamin, G., Paquot, M., Thonart, P., & Blecker, C. (2016).
Physical, functional and structural characterization of the cell wall fractions from
baker’s yeast Saccharomyces cerevisiae. Food Chemistry, 194, 1149–1155.
Burini, J. A., Eizaguirre, J. I., Loviso, C., & Libkind, D. (2021). Non-conventional yeasts
as tools for innovation and differentiation in brewing. Revista Argentina de
Microbiología, 53(4), 359–377.
Bzducha-Wróbel, A., Błażejak, S., Kawarska, A., Stasiak-Różańska, L., Gientka, I., &
Majewska, E. (2014). Evaluation of the efficiency of different disruption methods on
yeast cell wall preparation for β-glucan isolation. Molecules, 19(12), 20941–20961.
Carballo, C., Pinto, P. I., Mateus, A. P., Berbel, C., Guerreiro, C. C., Martinez-Blanch, J. F.,
et al. (2019). Yeast β-glucans and microalgal extracts modulate the immune response
and gut microbiome in Senegalese sole (Solea senegalensis). Fish & Shellfish
Immunology, 92, 31–39.
Caruso, A., Abraham, A., & Medrano, M. (2021, November). Alternative application for
β-glucans present in brewery yeast walls: Biological activity in vitro. In Poster session
presentation at the international workshop of brewing yeasts (IWOBY – IPATEC).
Bariloche, Argentina.
Cerletti, C., Esposito, S., & Iacoviello, L. (2021). Edible mushrooms and beta-glucans:
Impact on human health. Nutrients, 13(7). Article 2195.
Chaikliang, C., Wichienchot, S., Youravoug, W. Y., & Potchanapond, G. (2015).
Evaluation on prebiotic properties of B-glucan and oligo-B-glucan from mushrooms
by human faecal microbiota in fecal batch culture. Functional Foods in Health and
Disease, 5(11), 395–405.
Chaiyasut, C., Pengkumsri, N., Sivamaruthi, B. S., Sirilun, S., Kesika, P., Saelee, M., et al.
(2018). Extraction of β-glucan of Hericium erinaceus, Avena sativa L., and
Saccharomyces cerevisiae and in vivo evaluation of their immunomodulatory effects.
Food Science and Technology, 38, 138–146.
Ciecierska, A., Drywien, M., Hamulka, J., & Sadkowski, T. (2019). Nutraceutical
functions of beta-glucans in human nutrition. Roczniki Panstwowego Zakladu Higieny,
70(4), 315–324.
Cimini, A., & Moresi, M. (2021). Circular economy in the brewing chain. Italian Journal of
Food Science, 33(3), 47–69.
Da Silva Araújo, V. B., De Melo, A. N. F., Costa, A. G., Castro-Gomez, R. H.,
Madruga, M. S., Souza, E. L. De, et al. (2014). Followed extraction of β-glucan and
mannoprotein from spent brewer’s yeast (Saccharomyces uvarum) and application
of the obtained mannoprotein as a stabilizer in mayonnaise. Innovative Food Science
& Emerging Technologies, 23, 164–170. Elsevier.
De Marco Castro, E., Calder, P. C., & Roche, H. M. (2021). β-1, 3/1, 6-glucans and
immunity: State of the art and future directions. Molecular Nutrition & Food Research,
65(1). Article: 1901071.
De Vries, H., Geervliet, M., Jansen, C. A., Rutten, V. P., Van Hees, H., Groothuis, N., et al.
(2020). Impact of yeast-derived β-glucans on the porcine gut microbiota and immune
system in early life. Microorganisms, 8(10). Article 1573.
Deshpande, P. S., Sankh, S. N., & Arvindekar, A. U. (2011). Study of two pools of
glycogen in Saccharomyces cerevisiae and their role in fermentation performance.
Journal of the Institute of Brewing, 117(1), 113–119.
Dimopoulos, G., Tsantes, M., & Taoukis, P. (2020). Effect of high pressure
homogenization on the production of yeast extract via autolysis and beta-glucan
recovery. Innovative Food Science & Emerging Technologies, 62, Article 102340.
Elsevier.
Du, B., Bian, Z., & Xu, B. (2014). Skin health promotion effects of natural beta-glucan
derived from cereals and microorganisms: A review. Phytotherapy Research, 28(2),
159–166.
Farcas, A. C., Galanakis, C. M., Socaciu, C., Pop, O. L., Tibulca, D., Paucean, A., et al.
(2021). Food security during the pandemic and the importance of the bioeconomy in
the new era. Sustainability, 13(1), 150.
Ferreira, I., Pinho, O., Vieira, E., & Tavarela, J. G. (2010). Brewer’s Saccharomyces yeast
biomass: Characteristics and potential applications. Trends in Food Science &
Technology, 21(2), 77–84.
Freimund, S., Sauter, M., Käppeli, O., & Dutler, H. (2003). A new non-degrading isolation
process for 1, 3-β-D-glucan of high purity from baker’s yeast Saccharomyces
cerevisiae. Carbohydrate Polymers, 54(2), 159–171.
M.A. Caruso et al.
Fuller, R., Moore, M. V., Lewith, G., Stuart, B. L., Ormiston, R. V., Fisk, H. L., et al.
(2017). Yeast-derived β-1, 3/1, 6 glucan, upper respiratory tract infection and innate
immunity in older adults. Nutrition, 39, 30–35.
Geller, A., & Yan, J. (2020). Could the induction of trained immunity by β-glucan serve as
a defense against COVID-19? Frontiers in Immunology, 11. Article: 1782.
Geoghegan, I., Steinberg, G., & Gurr, S. (2017). The role of the fungal cell wall in the
infection of plants. Trends in Microbiology, 25(12), 957–967.
de Graaff, P., Berrevoets, C., Rӧsch, C., Schols, H. A., Verhoef, K., Wichers, H. J., et al.
(2021). Curdlan, zymosan and a yeast-derived β-glucan reshape tumor-associated
macrophages into producers of inflammatory chemo-attractants. Cancer Immunology,
Immunotherapy, 70(2), 547–561.
Gu, Y., Takagi, Y., Nakamura, T., Hasegawa, T., Suzuki, I., Oshima, M., et al. (2005).
Enhancement of radioprotection and anti-tumor immunity by yeast-derived β-glucan
in mice. Journal of Medicinal Food, 8(2), 154–158.
Hamza, Z., El-Hashash, M., Aly, S., Hathout, A., Soto, E., Sabry, B., et al. (2019).
Preparation and characterization of yeast cell wall beta-glucan encapsulated humic
acid nanoparticles as an enhanced aflatoxin B1 binder. Carbohydrate Polymers, 203,
185–192.
Helkar, P. B., Sahoo, A. K., & Patil, N. J. (2016). Food industry by-products used as a
functional food ingredient. International Journal of Wine Research, 6(3), 1–6.
Hofer, M., & Pospísi, l M. (1997). Glucan as stimulator of hematopoiesis in normal and
gamma-irradiated mice. A survey of the authors’ results. International Journal of
Immunopharmacology, 19(9–10), 607–609.
Hong, F., Hansen, R. D., Yan, J., Allendorf, D. J., Baran, J. T., Ostroff, G. R., et al. (2003).
Beta-glucan functions as an adjuvant for monoclonal antibody immunotherapy by
recruiting tumoricidal granulocytes as killer cells. Cancer Research, 63(24),
9023–9031.
Hromádková, Z., Ebringerová, A., Sasinková, V., Šandula, J., Hříbalová, V., &
Omelková, J. (2003). Influence of the drying method on the physical properties and
immunomodulatory activity of the particulate (1→ 3)-β-D-glucan from
Saccharomyces cerevisiae. Carbohydrate Polymers, 51(1), 9–15.
Ikewaki, N., Dedeepiya, V. D., Raghavan, K., Rao, K. S., Vaddi, S., Osawa, H., et al.
(2022). β-glucan vaccine adjuvant approach for cancer treatment through immune
enhancement (B-VACCIEN) in specific immunocompromised populations. Oncology
Reports, 47(1), 1–9.
Jacob, F. F., Striegel, L., Rychlik, M., Hutzler, M., & Methner, F. J. (2019a). Spent yeast
from brewing processes: A biodiverse starting material for yeast extract production.
Fermentation, 5(2), 51.
Jacob, F. F., Striegel, L., Rychlik, M., Hutzler, M., & Methner, F. J. (2019b). Yeast extract
production using spent yeast from beer manufacture: Influence of industrially
applicable disruption methods on selected substance groups with biotechnological
relevance. European Food Research and Technology, 245(6), 1169–1182.
Jaehrig, S. C., Rohn, S., Kroh, L. W., Wildenauer, F. X., Lisdat, F., Fleischer, L. G., et al.
(2008). Antioxidative activity of (1→ 3),(1→ 6)-β-d-glucan from Saccharomyces
cerevisiae grown on different media. LWT–Food Science and Technology, 41(5),
868–877.
Jagadiswaran, B., Alagarasan, V., Palanivelu, P., Theagarajan, R., Moses, J. A., &
Anandharamakrishnan, C. (2021). Valorization of food industry waste and by-
products using 3D printing: A study on the development of value-added functional
cookies. Future Foods, 4. Article 100036.
Kerby, C., & Vriesekoop, F. (2017). An overview of the utilisation of brewery by-products
as generated by british craft breweries. Beverages, 3(2). Article 24.
Khan, T., Date, A., Chawda, H., & Patel, K. (2019). Polysaccharides as potential
anticancer agents—a review of their progress. Carbohydrate Polymers, 210, 412–428.
Khan, A. A., Gani, A., Masoodi, F. A., Amin, F., Wani, I. A., Khanday, F. A., et al. (2016).
Structural, thermal, functional, antioxidant & antimicrobial properties of β-d-glucan
extracted from baker’s yeast (Saccharomyces cereviseae)—effect of γ-irradiation.
Carbohydrate Polymers, 140, 442–450.
Kittisuban, P., Ritthiruangdej, P., & Suphantharika, M. (2014). Optimization of
hydroxypropylmethylcellulose, yeast β-glucan, and whey protein levels based on
physical properties of gluten-free rice bread using response surface methodology.
LWT–Food Science and Technology, 57(2), 738–748.
Klis, F. M., Mol, P., Hellingwerf, K., & Brul, S. (2002). Dynamics of cell wall structure in
Saccharomyces cerevisiae. FEMS Microbiology Reviews, 26(3), 239–256.
Kogan, G., Staško, A., Bauerová, K., Polovka, M., Šoltés, L., Brezová, V., et al. (2005).
Antioxidant properties of yeast (1→ 3)-β-D-glucan studied by electron paramagnetic
resonance spectroscopy and its activity in the adjuvant arthritis. Carbohydrate
Polymers, 61(1), 18–28.
Kollár, R., Reinhold, B. B., Petráková, E., Yeh, H. J., Ashwell, G., Drgonová, J., et al.
(1997). Architecture of the yeast cell wall: β (1→ 6)-glucan interconnects
mannoprotein, β (1→ 3)-glucan, and chitin. Journal of Biological Chemistry, 272(28),
17762–17775.
Lam, K. L., & Cheung, P. C. K. (2013). Non-digestible long chain beta-glucans as novel
prebiotics. Bioactive Carbohydrates and Dietary Fibre, 2(1), 45–64.
Li, B., Allendorf, D. J., Hansen, R., Marroquin, J., Ding, C., Cramer, D. E., et al. (2006).
Yeast β-glucan amplifies phagocyte killing of iC3b-opsonized tumor cells via
complement receptor 3-Syk-phosphatidylinositol 3-kinase pathway. The Journal of
Immunology, 177(3), 1661–1669.
Li, B., Cai, Y., Qi, C., Hansen, R., Ding, C., Mitchell, T. C., et al. (2010). Orally
administered particulate β-glucan modulates tumor-capturing dendritic cells and
improves antitumor T-cell responses in cancer. Clinical Cancer Research, 16(21),
5153–5164.
Liepins, J., Kovačova, E., Shvirksts, K., Grube, M., Rapoport, A., & Kogan, G. (2015).
Drying enhances immunoactivity of spent brewer’s yeast cell wall β-d-glucans.
Journal of Biotechnology, 206, 12–16.
11
Food Hydrocolloids 133 (2022) 107963
Liu, D., Ding, L., Sun, J., Boussetta, N., & Vorobiev, E. (2016). Yeast cell disruption
strategies for recovery of intracellular bio-active compounds—a review. Innovative
Food Science & Emerging Technologies, 36, 181–192.
Liu, X. Y., Wang, Q., Cui, S. W., & Liu, H. Z. (2008). A new isolation method of β-d-
glucans from spent yeast Saccharomyces cerevisiae. Food Hydrocolloids, 22, 239–247.
Elsevier.
Lopes, L. M. F., de Mello, M. M. M., & Urbinati, E. C. (2022). β-Glucan reduces cortisol
plasma levels, enhances innate immune system after A. hydrophila inoculation, and
has lipolytic effects on the pacu (Piaractus mesopotamicus). Aquaculture, 546, Article
737411.
Magnani, M., Calliari, C. M., de Macedo, F. C., Mori, M. P., de Syllos Cólus, I. M., &
Castro-Gomez, R. J. H. (2009). Optimized methodology for extraction of (1 → 3)(1 →
6)-β-d-glucan from Saccharomyces cerevisiae and in vitro evaluation of the
cytotoxicity and genotoxicity of the corresponding carboxymethyl derivative.
Carbohydrate Polymers, 78, 658–665. Elsevier.
Majtan, J., & Jesenak, M. (2018). β-Glucans: Multi-functional modulator of wound
healing. Molecules, 23(4). Article 806.
Manners, D. J., & Meyer, M. T. (1977). The molecular structures of some glucans from
the cell walls of Schizosaccharomyces pombe. Carbohydrate Research, 57, 189–203.
Mantovani, M. S., Bellini, M. F., Angeli, J. P. F., Oliveira, R. J., Silva, A. F., &
Ribeiro, L. R. (2008). β-Glucans in promoting health: Prevention against mutation
and cancer. Mutation Research: Reviews in Mutation Research, 658(3), 154–161.
Marinescu, G., Stoicescu, A., & Patrascu, L. (2011). The preparation of mayonnaise
containing spent brewer’s yeast β-glucan as a fat replacer. Romanian Biotechnological
Letters, 16(2), 6017–6025.
Marson, G. V., de Castro, R. J. S., Belleville, M. P., & Hubinger, M. D. (2020). Spent
brewer’s yeast as a source of high added value molecules: A systematic review on its
characteristics, processing and potential applications. World Journal of Microbiology
and Biotechnology, 36(7), 1–22.
Martins, Z. E., Erben, M., Gallardo, A. E., Silva, R., Barbosa, I., Pinho, O., et al. (2015).
Effect of spent yeast fortification on physical parameters, volatiles and sensorial
characteristics of home-made bread. International Journal of Food Science and
Technology, 50(8), 1855–1863.
Medeiros, S. D. V., Cordeiro, S. L., Cavalcanti, J. E. C., Melchuna, K. M., Lima, A. M. D. S.,
Filho, I. A., et al. (2012). Effects of purified Saccharomyces cerevisiae (1→ 3)-β-glucan
on venous ulcer healing. International Journal of Molecular Sciences, 13(7),
8142–8158.
Medina-Gali, R., Ortega-Villaizán, M. M., Mercado, L., Novoa, B., Coll, J., & Perez, L.
(2018). Beta-glucan enhances the response to SVCV infection in zebrafish.
Developmental & Comparative Immunology, 84, 307–314.
Medrano, M., Gangoiti, M. V., Simonelli, N., & Abraham, A. G. (2020). Kefiran
fermentation by human faecal microbiota: Organic acids production and in vitro
biological activity. Bioactive Carbohydrates and Dietary Fibre, 24. Article: 100229.
Medrano, M., Simonelli, N., Miller, F., Piermaria, J., & Abraham, A. G. (2020,
November). Dietary fiber: Production of organic acids by the intestinal microbiota
and impact on cell proliferation: In vitro studies. In Poster session presentation at the
XIX meeting of the argentinian cancerology society. Buenos Aires, Argentina.
Middelberg, A. P. (1995). Process-scale disruption of microorganisms. Biotechnology
Advances, 13(3), 491–551.
Mo, L., Chen, Y., Li, W., Guo, S., Wang, X., An, H., et al. (2017). Anti-tumor effects of
(1→3)-β-d-glucan from Saccharomyces cerevisiae in S180 tumor-bearing mice.
International Journal of Biological Macromolecules, 95, 385–392.
Moretti, A. F., Moure, M. C., Quiñoy, F., Esposito, F., Simonelli, N., Medrano, M., et al.
(2022). Water Kefir, A fermented beverage containing probiotic microorganisms:
From ancient and artisanal manufacture to industrialized and regulated
commercialization. Future Foods. Article 100123.
Murphy, E. J., Rezoagli, E., Major, I., Rowan, N. J., & Laffey, J. G. (2020). β-glucan
metabolic and immunomodulatory properties and potential for clinical application.
Journal of Fungi, 6(4), 356.
Nakashima, A., Yamada, K., Iwata, O., Sugimoto, R., Atsuji, K., Ogawa, T., et al. (2018).
β-Glucan in foods and its physiological functions. Journal of Nutritional Science and
Vitaminology, 64(1), 8–17.
Natakankitkul, S. (2016). Development of skincare cosmetic from yeast beta-glucans.
Thai Journal of Pharmaceutical Sciences, 40, 9–12.
Novák, M., Synytsya, A., Gedeon, O., Slepička, P., Procházka, V., Synytsya, A., et al.
(2012). Yeast β (1-3),(1-6)-d-glucan films: Preparation and characterization of some
structural and physical properties. Carbohydrate Polymers, 87(4), 2496–2504.
Omara, I. I., Pender, C. M., White, M. B., & Dalloul, R. A. (2021). The modulating effect
of dietary beta-glucan supplementation on expression of immune response genes of
broilers during a coccidiosis challenge. Animals, 11(1). Article 159.
Øverland, M., Karlsson, A., Mydland, L. T., Romarheim, O. H., & Skrede, A. (2013).
Evaluation of Candida utilis, Kluyveromyces marxianus and Saccharomyces cerevisiae
yeasts as protein sources in diets for Atlantic salmon (Salmo salar). Aquaculture, 402,
1–7.
Pan, P., Huang, Y. W., Oshima, K., Yearsley, M., Zhang, J., Arnold, M., et al. (2019). The
immunomodulatory potential of natural compounds in tumor-bearing mice and
humans. Critical Reviews in Food Science and Nutrition, 59(6), 992–1007.
Patel, C. M., Chakraborty, M., & Murthy, Z. V. P. (2016). Fast and scalable preparation of
starch nanoparticles by stirred media milling. Advanced Powder Technology, 27(4),
1287–1294.
Peltzer, M. A., Salvay, A. G., Delgado, J. F., de la Osa, O., & Wagner, J. R. (2018). Use of
residual yeast cell wall for new biobased materials production: Effect of
plasticization on film properties. Food and Bioprocess Technology, 11(11), 1995–2007.
Pengkumsri, N., Sivamaruthi, B. S., Sirilun, S., Peerajan, S., Kesika, P., Chaiyasut, K.,
et al. (2016). Extraction of β-glucan from Saccharomyces cerevisiae: Comparison of
M.A. Caruso et al.
different extraction methods and in vivo assessment of immunomodulatory effect in
mice. Food Science and Technology, 37, 124–130.
Petravić-Tominac, V., Zechner-Krpan, V., Grba, S., Srečec, S., Panjkota-Krbavčić, I., &
Vidović, L. (2010). Biological effects of yeast β-glucans. Agriculturae Conspectus
Scientificus, 75(4), 149–158.
Piermaria, J., Rivero, S., & Medrano, M. (2021, August). Structural characterization by
ATR-FTIR of B-glucans obtained from walls of brewery yeasts. In Poster session
presentation at the meeting 6◦ Argentine symposium on biotechnological processes.
Misiones, Argentina.
Pinto, M., Coelho, E., Nunes, A., Brandão, T., & Coimbra, M. A. (2015). Valuation of
brewers spent yeast polysaccharides: A structural characterization approach.
Carbohydrate Polymers, 116, 215–222.
Piotrowska, A., Waszkiewicz-Robak, B., & Świderski, F. (2009). Possibility of β-glucan
from spent brewer’s yeast addition to yoghurts. Polish Journal of Food and Nutrition
Science, 59, 299–302.
Pizarro, S., Ronco, A. M., & Gotteland, M. (2014). ß-glucanos:¿qué tipos existen y cuáles
son sus beneficios en la salud? Revista Chilena de Nutricion, 41(4), 439–446.
Puligundla, P., Mok, C., & Park, S. (2020). Advances in the valorization of spent brewer’s
yeast. Innovative Food Science & Emerging Technologies, 62. Article 102350.
Rachwał, K., Waśko, A., Gustaw, K., & Polak-Berecka, M. (2020). Utilization of brewery
wastes in food industry. PeerJ, 8. Article e9427.
Raikos, V., Grant, S. B., Hayes, H., & Ranawana, V. (2018). Use of β-glucan from spent
brewer’s yeast as a thickener in skimmed yogurt: Physicochemical, textural, and
structural properties related to sensory perception. Journal of Dairy Science, 101(7),
5821–5831.
Rakowska, R., Sadowska, A., Dybkowska, E., & Swiderski, F. (2017). Spent yeast as
natural source of functional food additives. Roczniki Panstwowego Zakladu Higieny, 68
(2), 115–121.
Rieder, A., Ballance, S., Böcker, U., & Knutsen, S. (2018). Quantification of 1, 3-β-D-
glucan from yeast added as a functional ingredient to bread. Carbohydrate Polymers,
181, 34–42.
Sabu, C., Rejo, C., Kotta, S., & Pramod, K. (2018). Bioinspired and biomimetic systems for
advanced drug and gene delivery. Journal of Controlled Release, 287, 142–155.
Salgado, M., Rodríguez-Rojo, S., & Cocero, M. J. (2017). Barley and yeast β-glucans as
new emulsifier agents for the development of aqueous natural antifungal
formulations. Carbohydrate Polymers, 174, 1114–1120.
Samuelsen, A. B. C., Schrezenmeir, J., & Knutsen, S. H. (2014). Effects of orally
administered yeast-derived beta-glucans: A review. Molecular Nutrition & Food
Research, 58(1), 183–193.
Sceni, P. (2021). Ingredientes multicomponentes a partir de levadura. Estudio de sus
propiedades interfaciales y su aplicación en alimentos. (Tesis de doctorado).
Universidad Nacional de Quilmes, Bernal, Argentina. In Disponible en RIDAA-UNQ
Repositorio Institucional Digital de Acceso Abierto de la Universidad Nacional de Quilmes.
http://ridaa.unq.edu.ar/handle/20.500.11807/2947.
Sharma, R., Kataria, A., Sharma, S., & Singh, B. (2022). Structural characterisation,
biological activities and pharmacological potential of glycosaminoglycans and
oligosaccharides: A review. International Journal of Food Science and Technology, 57
(1), 4–15.
da Silva Guedes, J., Pimentel, T. C., Diniz-Silva, H. T., da Cruz Almeida, E. T.,
Tavares, J. F., de Souza, E. L., et al. (2019). Protective effects of β-glucan extracted
from spent brewer yeast during freeze-drying, storage and exposure to simulated
gastrointestinal conditions of probiotic lactobacilli. Lebensmittel-Wissenschaft und
-Technologie, 116. Article 108496.
Silva, W. D. O., & Morais, D. C. (2021). Transitioning to a circular economy in developing
countries: A collaborative approach for sharing responsibilities in solid waste
management of a Brazilian craft brewery. Journal of Cleaner Production, 319. Article
128703.
Simonelli, N., Gagliarini, N., Medrano, M., Piermaria, J. A., & Abraham, A. G. (2020).
Kefiran. Ch 1 in: Polysaccharides of microbial origin: Biomedical applications. In
J. Oliveira, H. Radhouani, & R. L. Reis (Eds.), Polysaccharides of microbial origin (pp.
1–19). Cham. Oxford, United Kingdom: Springer.
Sivieri, K., de Oliveira, S. M., de Souza Marquez, A., Pérez-Jiménez, J., & Diniz, S. N.
(2022). Insights on β-glucan as a prebiotic coadjuvant in the treatment of diabetes
mellitus: A review. Food Hydrocolloids for Health. Article: 100056.
Sugawara, T., Takahashi, S., Osumi, M., & Ohno, N. (2004). Refinement of the structures
of cell-wall glucans of Schizosaccharomyces pombe by chemical modification and
NMR spectroscopy. Carbohydrate Research, 339(13), 2255–2265.
Suphantharika, M., Khunrae, P., Thanardkit, P., & Verduyn, C. (2003). Preparation of
spent brewer’s yeast β-glucans with a potential application as an immunostimulant
for black tiger shrimp, Penaeus monodon. Bioresource Technology, 88(1), 55–60.
Suwanapong, S., Khongsay, N., Laopaiboon, L., Jaisil, P., & Laopaiboon, P. (2013). Dried
spent yeast and its hydrolysate as nitrogen supplements for single batch and
repeated-batch ethanol fermentation from sweet sorghum juice. Energies, 6(3),
1618–1631.
Tamura, K., Hemsworth, G. R., Déjean, G., Rogers, T. E., Pudlo, N. A., Urs, K., et al.
(2017). Molecular mechanism by which prominent human gut Bacteroidetes utilize
12
Food Hydrocolloids 133 (2022) 107963
mixed-linkage beta-glucans, major health-promoting cereal polysaccharides. Cell
Reports, 21(2), 417–430.
Teferra, T. F. (2021). Possible actions of inulin as prebiotic polysaccharide: A review.
Food Frontiers, 2(4), 407–416.
Thammakiti, S., Suphantharika, M., Phaesuwan, T., & Verduyn, C. (2004). Preparation of
spent brewer’s yeast β-glucans for potential applications in the food industry.
International Journal of Food Science and Technology, 39(1), 21–29.
Thanardkit, P., Khunrae, P., Suphantharika, M., & Verduyn, C. (2002). Glucan from spent
brewer’s yeast: Preparation, analysis and use as a potential immunostimulant in
shrimp feed. World Journal of Microbiology and Biotechnology, 18(6), 527–539.
Thomas, S., Rezoagli, E., Abidin, I. Z., Major, I., Murray, P., & Murphy, E. J. (2022).
β-Glucans from yeast—immunomodulators from novel waste Resources. Applied
Sciences, 12(10), 5208.
Tian, X., Shao, Y., Wang, Z., & Guo, Y. (2016). Effects of dietary yeast β-glucans
supplementation on growth performance, gut morphology, intestinal Clostridium
perfringens population and immune response of broiler chickens challenged with
necrotic enteritis. Animal Feed Science and Technology, 215, 144–155.
Vetvicka, V., & Vetvickova, J. (2011). β (1-3)-D-glucan affects adipogenesis, wound
healing and inflammation. Oriental Pharmacy and Experimental Medicine, 11(3),
169–175.
Vetvicka, V., & Vetvickova, J. (2018). Glucans and cancer: Comparison of commercially
available β-glucans–Part IV. Anticancer Research, 38(3), 1327–1333.
Wagner, J. R., Sceni, P., & Otero Rambla, M. A. (2008). Polisacáridos Estructurales de
levadura. Glucanos y mananos. In J. R. Wagner, M. A. Otero Rambla, & I. Guerrero
Legarreta (Eds.), Las Levaduras y sus productos derivados como ingredientes en la
industria de los alimentos (pp. 233–246). Bernal: Universidad Nacional de Quilmes.
Wang, H., Chen, G., Li, X., Zheng, F., & Zeng, X. (2020). Yeast β-glucan, a potential
prebiotic, showed a similar probiotic activity to inulin. Food & Function, 11,
10386–10396.
Wang, N., Liu, H., Liu, G., Li, M., He, X., Yin, C., et al. (2020). Yeast β-D-glucan exerts
antitumour activity in liver cancer through impairing autophagy and lysosomal
function, promoting reactive oxygen species production and apoptosis. Redox
Biology, 32. Article: 101495.
Waszkiewicz-Robak, B., & Bartnikowska, E. (2009). Effects of spent brewer’s yeast and
biological β-glucans on selected parameters of lipid metabolism in blood and liver in
rats. Journal of Animal and Feed Sciences, 18(4), 699–708.
Watanabe, N., Inaida, O., Yamada, K., & Karakawa, T. (1980). New approaches to using
spent brewer’s yeast. Journal of the American Society of Brewing Chemists, 38(1), 5–8.
Worrasinchai, S., Suphantharika, M., Pinjai, S., & Jamnong, P. (2006). β-Glucan prepared
from spent brewer’s yeast as a fat replacer in mayonnaise. Food Hydrocolloids, 20(1),
68–78.
Xavier-Santos, D., Padilha, M., Fabiano, G. A., Vinderola, G., da Cruz, A. G., Sivieri, K.,
et al. (2022). Evidences and perspectives of the use of probiotics, prebiotics,
synbiotics, and postbiotics as adjuvants for prevention and treatment of COVID-19: A
bibliometric analysis and systematic review. Trends in Food Science & Technology,
120, 174–192.
Yiannikouris, A., Francois, J. E. A. N., Poughon, L., Dussap, C. G., Bertin, G., Jeminet, G.,
et al. (2004). Adsorption of zearalenone by β-D-glucans in the Saccharomyces
cerevisiae cell wall. Journal of Food Protection, 67(6), 1195–1200.
Yoo, M. S., & Lee, Y. T. (2007). Pasting properties of crude beta-glucan from spent
brewer’s yeast on wheat flour and starch. Food Science and Biotechnology, 16(3),
485–488.
Yurdacan, H. M., & Sari, M. M. (2021). Functional green-based nanomaterials towards
sustainable carbon capture and sequestration. In J. Hayton (Ed.), Sustainable
materials for transitional and alternative energy (pp. 125–177). Oxford. United
Kingdom: Gulf Professional Publishing (Elsevier). Ch 3 in.
Zhong, K., Liu, Z., Lu, Y., & Xu, X. (2021). Effects of yeast β-glucans for the prevention
and treatment of upper respiratory tract infection in healthy subjects: A systematic
review and meta-analysis. European Journal of Nutrition, 1–13.
Zhu, F., Du, B., & Xu, B. (2016). A critical review on production and industrial
applications of beta-glucans. Food Hydrocolloids, 52, 275–288.
Web references
Reference 1: Retrieved from: https://www.statista.com/statistics/270275/worldwi
de-beer-production/Accessed February 17, 2022.
Reference 2: Retrieved from https://www.cfsanappsexternal.fda.gov/scripts/fdcc/?se
t=GRASNotices&id=239&sort=GRN_No&order=DESC&startrow=1&type=ba
sic&search=glucan Accessed February 17, 2022.
Reference 3: Retrieved from http://www.fda.gov/Food/IngredientsPackagingLabeling/
GRAS/NoticeInventory/ucm153925.htm Accessed February 17, 2022.
Reference 4: Retrieved from https://www.wellmune.com/Accessed February 17, 2022.
Reference 5: Retrieved from: https://www.wellmune.com/2011/03/07/biotheras-well
mune-wgp-approved-for-food-use-in-china/?lang=es Accessed February 17, 2022.
Reference 6: retrieved from: http://www.conal.gob.ar/Accessed February 17, 2022.